AN ABSTRACT OF THE THESIS OF

Chun -shu Lee for the Master of Science (Name of student) (Degree) in Botany presented on iq 6v (Major) ih ( Dal %)

Title: COMPARATIVE WOOD ANATOMY OF THE FAGACEAE

OF TAIWAN Abstract approved Frank H, Smith

Many species of the Fagaceae have been variously interpre- ted on the basis, primarily of external morphological characteristics.

The objectives of this study are to elucidate generic and specific characteristics of the wood of Fagaceae and to attempt to obtain ev- idence regarding taxonomic relationships among some of the species occurring in Taiwan that have been variously interpreted.

There are 45 species (Lin and Liu, 1965) of Fagaceae, sub- divided to seven genera, in Taiwan and 35 of these were obtained for study. Detailed descriptions are provided for all species and dif- ferences among species and genera are emphasized.

On the basis of comparative wood anatomy of the species studied, several suggestions are made regarding relationships of certain species.

The wood anatomy of Pasania uraiana (Hay.) Schottky is very similar to that of Castanopsis and quite different from the anatomy

of other species of Pasania or of Lithocarpus. The cupula is cup- like, however, so that it differs in this important generic character from Castanopsis. It probably should be treated under a distinct genus, Limlia, as Masamune and Tomiya (1947) suggested.

The comparison of wood anatomy between Pasania dodoniae- folia and Pasania formosana shows consistent differences. Pasania dodoniaefolia shows distinct growth rings, homogeneous rays and absence of crystals in the parenchyma cells. Pasania formosana

shows indistinct growth rings, heterogeneous rays and presence of crystals in parenchyma cells. Since the morphological characters are also somewhat different (Lin and Liu, 1965), they probably should be treated as two distinct species. Morphologically, Castanopsis carlesii var. sessilis is very similar to Castanopsis carlesii, but is quite different with regard to wood anatomy. Since Castanopsis carlesii var. sessilis has ring porous wood, conspicuous growth rings and pore size transition abrupt, while Castanopsis carlesii has diffuse porous wood, incon- spicuous growth rings, pore size transition gradual, these should be considered as separate species and the original name, Castanopsis

stipitata should be recognized as the valid name. The genera Lithocarpus and Pasania could not be separated anatomically. These genera have been separated primarily on the basis of the cupula but they are very similar with regard to other morphological characteristics. If both anatomy and external mor-

phology are considered, these should be treated as one genus, Litho - carpus. Comparative Wood Anatomy of the Fagaceae of Taiwan.

by

Chun -shu Lee

A. THESIS

submitted to Oregon State University

in partial fulfillment of the requirements for the degree of Master of Science June 1968 APPROVED

Professor of Botany In Charge of Major

Head of Department of otany and Plant Pathology

Dean of Graduate School

Date thesis is presented (f Typed by Ruth Baines ACKNOWLEDGMENTS

I want to express my deepest gratitude to Dr. Frank H. Smith,

Department of Botany, Oregon State University, who directed this thesis. I am especially indebted to him for his kind encouragement and careful guidance and for his help in the preparation of this thesis.

The personal interest and encouragement of my friend, Mr. Tsing Liu, Taxonomist, Taiwan Forestry Research Institute, Re- public of China, who at present is studying in the Department of Forest Management, Oregon State University, are most gratefully acknowledged. He provided the basic elements of the problem and counseled in taxonomical questions which arose.

I am grateful to my colleague, Mr. David Hansen, Department

of Botany and Plant Pathology, Oregon State University, for his help in the photographic aspects of this study.

My academic training and this study would not be achieved without a grant from the National Science Council, Republic of China.

Here I express my heartfelt gratefulness. TABLE OF CONTENTS

Page

INTRODUCTION 1

MATERIALS AND METHODS 4

SPECIES DESCRIPTIONS OF WOODS 5

ANATOMICAL KEY TO SPECIES OF FAGACEAE IN TAIWAN 55

DISCUSSION AND CONCLUSIONS 59 Fagus (Tourn.) Linn. 60 Castanea Tourn. 60 Castanopsis Spach. 61 Lithocarpus Bi . and Pasania Oerst 61 Quercus Linn. 62 Cyclobalanopsis Oerst 62

BIBLIOGRAPHY 69

APPENDIX 71 LIST OF TABLE

Table Page

1 Species of Fagaceae collected in Taiwan. 3 COMPARATIVE WOOD ANATOMY OF THE FAGACEAE OF TAIWAN

INTRODUCTION

Trees of the Fagaceae are the principal components of the broad - leaved forests to Taiwan. Because of the large size and com- mercial value of the wood, they are considered to be economically important for use as lumber.

Originally, only five genera of the Fagaceae were recognized in Taiwan, They are Fagus (Linn. Syst. ed. I, 1735), Castanea

(Tourn. ex Linn. Syst. ed, I, 1735), Gastanopsis (Spach. Hist. Veg.

Phan. XI, 185, 1842), Lithocarpus (Blume, Bijdr. 526, 1825) and

Quercus (Linn. Syst. ed. I, 1735). In more recent years, four ad- ditional genera have been recognized by various authors. They are

Pasania (Oerst. in Kjoeb. Vidensk. Med. 81, 1866), Cyclobalanopsis

(Oerst. in Kjoeb. Vidensk. Med. 77, 1866), Shiia (Makino in Journ.

Jap. Bot. V, 23, 1928) and Limlia (Masam. et Tomiya in Acta Bot.

Taiwan, I, 1, 1947). Actually, Pasania was separated from Litho- carpus; Cyclobalanopsis was separated from Quercus; Shiia and

Limlia were derived from some species of Castanopsis and were recognized by only a few Japanese taxonomists.

A review of the systematic studies of the Fagaceae of Taiwan shows that several species have been interpreted differently by various authors. The problematic species which have been 2 considered by the author in this study are Castanopsis carlesii

(Hems'. ) Hay. var. sessilis Nakai, which was treated as a distinct species, Castanopsis stipitata by Kanehira (1939); Pasania uraiana (Hay.) Schottky, which has been treated under various genera by var- ious taxonomists because of the confusing characters of the plant; Pasania formosana (Skan) Schottky and Pasania dodoniaefolia Hay. which are considered to be conspectific by Li (1953), but Lin and Liu

(1965) deem that they are distinct species as originally described.

The objectives of this study are to elucidate generic and spe- cific characteristics of the wood of Fagaceae and to obtain evidence regarding taxonomic relationships among the species occurring in Taiwan.

According to Lin and Liu (1965), there are 45 species, sub- species and varieties of Fagaceae in Taiwan, among which are six introduced species. Unfortunately our wood collection is incomplete to date, but 35 species have been collected, of which four are intro- duced (Table 1). However, several of the species which have been variously interpreted are included, and it is hoped that this study might be helpful for solving some of the confusing classification problems. 3

Table 1. Species of Fagaceae collected in Taiwan. Introduced species are indicated by an asterisk

No. Scientific Name Collecting Date Place Collector

1 Fagus hayatae Palib. May 20, 1966 Mt. Nan- cha -tein C.N. Wu *2 Castanea creneta Sieb. et Zucc. Feb. 20. 1966 Liu -kuei C. S. Lee *3 Castanea mollissima Bl. Oct. 25, 1966 Wu -she C. S. Lee 4 Castanopsis carlesii (Hemsl. ) Hay. Feb. 19, 1966 Liu -kuei C. S. Lee 5 Castanopsis carlesii (Hemsl. ) Hay. var. sessilis Nak. Nov. 1, 1965 Lien -hua -chih C. S. Lee 6 Castanopsis subacuminata Hay. Feb. 18, 1966 Liu-kuei C. S. Lee 7 Castanopsis formosana Hay. Feb. 22, 1966 Liu-kuei C. S. Lee 8 Castanopsis hystrix A. DC. Nov. 2, 1965 Lien- hua -chih C.S. Lee 9 Castanopsis kawakamii Hay. Nov. 2, 1965 Lien- hua -chih C. S. Lee 10 Castanopsis kusanoi Hay. Feb. 23, 1966 Liu -kuei C. S. Lee 11 Castanopsis stellatospina Hay. Feb. 15, 1966 Mt. Nan -j en C. N. Chang 12 Lithocarpus amygdalifolius (Skan) Hay. Nov. 3, 1965 Lien- hua -chih C. S. Lee 13 Lithocarpus castanopsisifglius Hay. Feb. 25, 1966 Liu-kuei C.S. Lee 14 Pasania kodaihoensis (Hay. ) H. L. Li Feb. 23', 1966 Liu -kuei C. S. Lee 15 Pasania konishii (Hay. ) Schottky Nov. 4, 1965 Lien- hua -chih C. S. Lee 16 Pasania brevicaudata (Skan) Schottky Nov. 1, 1965 Lien- hua -chih C. S. Lee 17 Pasania uraiana,(Hay.) Schottky Nov. 3, 1965 Lien -hua -chih C. S. Lee 18 Pasania kawakamii (Hay. ) Schottky Feb. 26, 1966 Liu -kuei C. S. Lee 19 Pasania dodoniaefolia Hay. Apr. 20, 1966 Taitung C.N. Chang 20 Pasania randaiensis (Hay. ) Schottky Nov. 5, 1965 Lien- hua -chih C. S. Lee 21 Pasania formosana (Skan) Schottky Feb. 15, 1966 Mt. Nan -j en C.N. Chang 22 Pasania ternaticupula (Hay. ) Schottky Nov. 4, 1965 Lien- hua -chih C. S. Lee 23 Pasania nantoensis (Hay. ) Schottky Nov. 5, 1965 Li en- hua -chih C. S. Lee 24 Quercus variabilis Bl. Sept. 10, 1965 Taipei C.S. Lee *25 Quercus acutissima Canna, Sept. 10, 1965 Taipei C. S. Lee *26 Quercus phillyraeoides A. Gray Sept. 11, 1965 Taipei C. S. Lee 27 Quercus semicarpifolia Smith Subsp. glabra (Franch. ) Handel -Matz. Oct. 9, 1966 Li Shan T. Liu 28 Cyclobalanopsis pachyloma (O. Seem. ) Schottky Nov. 13, 1965 Lien- hua -chih C.S. Lee 29 Cyclobalanopsis morii (Hay. ) Schottky May 6, 1966 Mt. Ta -hsueh C.S. Lee 30 Cyclobalanopsis championii (Benth. ) Oerst. Feb. 13, 1966 Mt. Nan -j en C. N. Yang 31 Cyclobalanopsis gilva (BL ) Oerst. Sept. 9, 1965 Taipei C.S. Lee 32 Cyclobalanopsis stenophyiloides (Hay. ) Kudo et Mas am. Feb. 24, 1966 Liu -kuei C.S. Lee 33 Cyclobalanopsis glauca (Thunb.) Oerst. Sept. 19, 1965 Taipei C.S. Lee 34 Cyclobalanopsis longinux (Hay.) Schottky Feb. 26, 1966 Liu -kuei C.S. Lee 35 Cyclobalanopsis globosa Lin et Liu Sept. 18, 1966 Kuan -tao -hsi Y.C. Liu 4

MATERIALS AND METHODS

All wood samples were collected from stems or branches at

least 3 cm in diameter. Three blocks, 1x1x2 cm were cut from each

sample so as to obtain cross, radial and tangential sections of each

species. Since wood of the Fagaceae is very hard and difficult to cut with a microtome, all blocks were softened in 25% hydrofluoric acid for 6 -8 weeks before sectioning. Thickness of sections varied with

species and/or plane of sectioning. Sections were stained with Del- afield' s haematoxylin and safranin O and prepared as permanent slide specimens. Slides were also prepared from wood macerated accord-

ing to the method of Jeffery (Johansen, 1940).

The wood of each species was analyzed with regard to the following characteristics:

(a) Distinctness of annual rings, and difference between early wood and late wood.

(b) Distribution, morphology, perforation, structure, and

patterns of vessels and tracheids,

(c) Distribution and morphology of fibers.

(d) Distribution and structure of xylem paranchyma strands.

(e) Distribution, structure and cell types of rays. Cross, radial and tangential sections were photographed

separately, and reproduced at the same magnification so that direct comparisons can be made among the different species. 5

SPECIES DESCRIPTIONS OF WOODS

1. Fagus hayatae Palib in Hay. Mater. Fl. Formos. (1911) 286.

Wood diffuse porous; growth ring distinct, delineated by a band of denser late wood.

Pores evenly distributed, those in the early wood usually some- what larger but the transition in pore size from early to late wood gradual, 150 -225 per mm2; solitary and aggregated in clusters, tan- gential or radial rows of 2- several, polygonal, thin -walled, 1-1.5 µ in wall thickness, 15 -61 p. in diameter; end walls oblique, perforation plates simple, reticulate and scalariform, 12 -25 bars; pits leading to contiguous tracheal cells bordered, pit cavity orbicular or ellip- tical, aperture lenticular; vessel members 138 -451 µ in length. Tyloses present in heartwood.

Fibers 3 -5 µ in wall thickness, 9 -19 µ in diameter, 467-1,018 p. in length.

Parenchyma apotracheal diffuse and paratracheal; (a) apotrach- eal diffuse parenchyma conspicuous, few cells with tannin inclusions;

(b) paratracheal parenchyma sparse, restricted to occasional cells

(not forming a sheath). Rays .nonstoried, homogeneous or with marginal upright cells, broad and narrow; (a) broad rays several- many- seriate, up to

269 µ in width, a few cells with tannin inclusions; (b) narrow rays 6

1 -3- seriate, 1 -23 cells in height.

2. Castanea crenata Sieb. et Zucc. in Abh. Akad. Muench. IV 3.

(1846), 324.

Syn. Castanea stricta Sieb. et Zucc. in Abh. Akad. Muench.

(1846) .225.

Castanea japonica Blume in Mus. Bot. Lugd. Bot. I (1849)

. 284.

Wood ring porous; growth ring conspicuous, delineated by a narrow band of flattened fibers.

Pores unevenly distributed; early wood pores very large forming a broad, conspicuous band several pores in width, 9 -21 per mm2, solitary or occasionally in short radial rows, in late wood aggregated, orbicular or elliptical, 2 -3 µ in wall thickness, 60 -216 µ in diameter; late wood pores small, arranged in flame -like patches, 55 -119 per mm2; the transition in pore size from early to late wood abrupt; end walls oblique, perforation plates simple or those in late wood occas- ionally scalariform, 3 -5 bars; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture lenticular; vessel members 181 -433 p. in length. Tyloses present in heartwood. Vas - icentric tracheids present, confined to the vicinity of the early wood vessels.

Fibers abundant in late wood, 3 -6 in wall thickness, 7 -12 µ 7 in diameter, 632-1,120 in length.

Parenchyma diffuse -in- aggregate and paratracheal; (a) diffuse - in- aggregate parenchyma abundant, especially in late wood; (b) para- tracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); a few cells with tannin inclusions. Rays nonstoried, homogeneous, narrow, uniseriate or rarely in part biseriate, 1 -19 cells in height.

3. Castanea mollissima Blume in Mus. Bot. Ludg. Bot. I (1849)

286.

Syn. Castanea bungeana Bl. in Mus. Bot. Lugd. Bot. I (1849)

284.

Castanea crenata sensu Henry in Elwer and Henry Trees

Great Brit. and Irel. IV (1909) 854. (pro parte) non S.

et Z. Castanea sativa Mill. var. mollissima Pampanini in Nuov.

Giron. Bot. Ital. n. ser. XVII (1910) 250. Castanea sativa Mill. var. formosana Hay. Mater. Fl.

Formos. (1911) 304.

Castanea formosana Hay. Gen. Ind. Fl. Formos. (1917) 71.

Castanea hupehensis Dode in Bull. Soc. Dendr. France

(1908) 151.

Wood ring porous; growth ring conspicuous, delineated by a 8 a narrow band of flattened fibers contiguous to a band of large pores.

Pores unevenly distributed; early wood pores very large form- ing a broad, conspicuous band several pores in width, 18 -30 per mm2, solitary, orbicular or elliptical, 2 -3 µ.in wall thickness, 49 -218 µ in diameter; late wood pores small, polygonal, arranged in flame -like patches, 172 -351 per mm2; the transition in pore size from early to late wood abrupt; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture elliptical; vessel members 95 -382 µ in length. Tyloses present in heartwood. Vasicentric tracheids present, sparse, con- fined to the vicinity of the early wood vessels.

Fibers arranged in radial rows, thin -walled, 1.5-3 p. in wall thickness, 5 -18 µ in diameter, 675 -965 µ in length.

Parenchyma diffuse -in- aggregate and paratracheal (a) diffuse - in- aggregate parenchyma abundant, very conspicuous, especially in late wood; (b) paratracheal parenchyma sparse, restricted to occas- ional cells (not forming a sheath); most of the cells with tannin in- clusions. Rays nonstoried, homogeneous, narrow, uniseriate or rarely in part biseriate, 2 -16 cells in height; most of the cells with tannin inclusions. 9

4. Castanopsis carlesii (Hemsl. ) Hay. Gen. Ind. Fl. Formos.

(1917) 72.

Syn. Castanopsis longicaudata (Hay.) Kaneh. et Hatus. in Trans.

Nat. Hist. Soc. Formos. XXX (1939) 155.

Quercus carlesii Hemsl. in Hook. Icon. pl. XXVI pt.

2591 (1899).

Pasania cuspidata sensu Mats. et Hay. Enum. Pl. Formos.

(1906) 392 non Oerst,

Quercus longicaudata Hay. Icon. pl. Formos. IV (1913) 182.

Pasania longicaudata Hay. 1. c.

Lithocarpus longicaudata Hay. Gen. Ind. Fl. Formos.

(1917) 72.

Shiia longicaudata Kudo et Masam. in Journ. Soc. Trop.

Agr. Formos. II (1930) 35.

Shiia carlesii (Hemsl. ) Kudo in Journ. Soc. Trap. Agr.

Fromos. III (1931) 17.

Synaedrys carlesii Koidz. in Bot. Mag. Tokyo XXX (1916)

186.

Wood diffuse porous; growth ring inconspicuous. Pores un- evenly distributed, arranged in stream -like clusters which extend for some distance radially across several to many rings; those in the early wood somewhat larger but the transition in pore size from early to late wood gradual; 11 -25 per mm2; solitary or in radial 10 groups of two, polygonal, 2 -3 p. in wall thickness, 28 -108 µin dia- meter; end walls oblique, perforation plates simple, very few scalariform with 1 -2 bars; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture elliptical; vessel members

175 -525 p. in length. Very few tyloses in heartwood. Vasicentric tracheids present, confined to the vicinity of larger vessels,

Fibers abundant, 2-3.5 µ in wall thickness, 7 -21 µ. in dia- meter, 610-1,120 in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma conspicuous, in numerous, concentric, continuous

1- 4- seriate (mostly 1 -2) ragged lines; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath).

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several -many- seriate, up to 124 µ in width, many cells (into the hundreds) up to 3,421 p. in height; aggregate rays composed of smaller rays separated by strands of fibrous tissue, otherwise com- parable to the compounds rays; (b) narrow rays very numerous, uniseriate, 1 -23 cells in height; a few cells with tannin inclusions.

5. Castanopsis carlesii (Hemsl. ) Hay. var. sessilis Nak. in Journ.

Jap, Bot. XV (1939) 261. 11

Syn. Castanopsis stipitata (Hay. ) Kaneh. in Trans. Nat. Hist.

Soc. Formos. XXX (1939) 155.

Quercus junghuhnii sensu Hay. Fl. Mont. Formos. (908)

203, non Miq.

Synaedrys stipitata Koidz. in Bot. Mag. Tokyo XXX

(1916) 187.

Quercus stipitata Hay. ex. Koidz. in Bot. Mag. Tokyo

XXX (1916) 187.

Lithocarpus stipitata Koidz. in 1. c. XXXIX (1925) 2.

Shiia stipitata Kudo et Masam. in 1. c. XLIV (1930) 406.

Quercus carlesii sensu Hay. Mater. F1. Formos. (1911)

289 non Hemsl.

Wood ring porous; growth ring conspicuous, delineated by a narrow band of flattened fibers contiguous to a broad band of large pores.

Pores unevenly distributed; early wood pores very large, forming a broad, conspicuous band several pores in width, 6 -11 per mm2, solitary or occasionally in pairs, polygonal, orbicular or elliptical, 3 -5 µ in wall thickness, 55 -177 µ in diameter; late wood pores small, polygonal, arranged in flame -like patches, 101 -153 per mm2; the transition in pore size from early to late wood abrupt; 12

end walls oblique, perforation plates simple or those in the smaller

late wood vessels occasionally scalariform, 1 -3 bars; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aper- ture orbicular or elliptical; vessel members 211 -635 µ in length.

Tyloses present in heartwood. Vasicentric tracheids present, con- fined to the vicinity of the early wood large vessels.

Fibers 2 -5 µ in wall thickness, 7 -22 H. in diameter, 568-

1, 00B p. in length.

Parenchyma apotracheal banded and paratracheal; (a) apo- tracheal banded parenchyma in numerous, concentric, continuous,

1- 3- seriate ragged tangential bands; (b) paratracheal parenchyma

sparse, restricted to occasional cells.

Rays nonstoried, heterogeneous; (a) aggregate rays com- posed of smaller rays separated by strands of fibrous tissue; (b) narrow rays very numerous, uniseriate or biseriate, 1 -25 cells in height; very few cells with tannin inclusions.

6. Castanopsis subacuminata Hay. Icon. Pl. Formos. III (1913)

189.

Syn. Castanopsis indica sensu Hay. F1. Mont. Formos. (1908)

204 non A. DC. 13

Wood diffuse porous; growth ring distinct in heartwood, indistinct in sap wood.

Pores unevenly distributed, arranged in stream -like clusters which extend for some distance radially, those in the early wood usually somewhat larger but the transition in pore size from the early to late wood gradual, 1 -11 per mm2; solitary, polygonal or oval, 2 -3 p. in wall thickness, 45 -227 p. in diameter; end wall oblique, perforation plates orbicular or elliptical, aperture lenticular; ves- sel members 211 -375 µ in length. Tyloses present in heartwood.

Vasicentric tracheids present, forming a 1 -3 seriate sheath, inter- rupted by parenchyma, about the vessels.

Fibers abundant, 3 -4 µ in wall thickness, 10 -75 µ in dia- meter, 623 -961 µ in length.

Parenchyma diffuse -in- aggregate and paratracheal; (a) diffuse -in- aggregate parenchyma abundant, a few cells with book - shaped crystals; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath).

Rays nonstoried, heterogeneous, narrow, uniseriate or rarely in part biseriate, 1 -21 cells in height. 14

7. Castanopsis formosana Hay. Icon. Pl. Formos. V (1913) 189.

Syn. Castanopsis tribuloides D. C. var. formosana Skan in

Journ. Linn. Soc. Bot. XXIV (1899) 524.

Wood diffuse porous; growth ring distinct, delineated by a band of thick -walled fibers.

Pores unevenly distributed, arranged in flame -like or stream -like clusters which extend for some distance radially, 6 -18 per mm2; solitary, polygonal, orbicular or oval, 2 -3 p. in wall thickness, 46 -139 p. in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular or elliptical, aperture lenticular; vessel mem- bers 139 -486 p. in length. Tyloses present in heartwood. Vasi- centric tracheids present, forming a sheath, interrupted by paren- chyma, about the vessels.

Fibers thick -walled, 3 -6 p. in wall thickness, 7 -15 p. in dia- meter, 628 -945 p. in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma abundant, in numerous, concentric, continuous, 1 -3- seriate ragged tangential bands; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); a few cells with 15 book- shaped crystals.

Rays nonstoried, heterogeneous or nearly so; uniseriate,

1 -20 cells in height; many cells with tannin inclusions.

8. Castanopsis hystrix A. DC. in Journ. Bot. (1863) 182.

Syn. Castanopsis taiwaniana Hay. F1. Mont. Formos. (1908)

205; Mater. F1. Formos. (1911) 303.

Wood ring porous; growth ring conspicuous, delineated by a narrow band of flattened fibers bordered with a broad band of large pores.

Pores unevenly distributed, early wood pores very large forming a conspicuous band several pores in width, 5 -12 per mm2,

solitary, polygonal, orbicular or oval, 204 N, in wall thickness, 61-

188 p. in diameter; late wood pores small, polygonal, arranged in flame- shaped clusters, 119 -197 per mm2, the transition in pore size from early to late wood abrupt; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular or elliptical, aperture lenticular; vessel mem- bers 172 -562 µ in length. Few tyloses present in heartwood. Vasi- centric tracheids present, forming 1- 3- seriate sheath, interrupted by parenchyma, confined to the vicinity of the early wood large 16 vessels.

Fibers arranged in numerous, concentric bands, 3 -6 p, in wall thickness, 8 -19 µin diameter, 553-1,126 µ in length. Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma abundant, conspicuous, in numerous, concentric, con- tinuous, 1- 3- seriate ragged tangential bands; (b) paratracheal par - encyma sparse, restricted to occasional cells.

Rays nonstoried, heterogeneous (a) compound and aggregate rays several- many- seriate, up to 133 µ in width, many cells, up to

1,672 p, in height; aggregate rays composed of smaller rays separat- ed by strands of fibrous tissue; (b) narrow rays numerous, uniseri- ate, 2-21 cells in height.

9. Castanopsis kawakamii Hay. Mater. Fl. Formos. (1911) 302.

Wood ring porous; growth ring conspicuous, delineated by a very narrow band of flattened fibers bordered with a 1- 3- seriate band of large vessels.

Pores unevenly distributed, early wood pores very large forming a conspicuous, tangential band l -3 pores in width, 6 -13 per mm2, solitary, orbicular or oval, 2 -4 µ in wall thickness, 49 -151 µ in diameter; late wood pores small, polygonal, arranged in broad, flame- shaped patches, 101 -188 per mm2, the transition in pores size from early to late wood abrupt; end walls oblique, perforation 17 plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular or elliptical, aperture elliptical or lenticular; vessel members 120 -485 µ in length. Tyloses occasionally present in vessels. Vasicentric tracheids forming a sheath interrupted by parenchyma, confined to the vicinity of the early wood large vessels. Fibers thick -walled, arranged in obliquely radial patches,

3 -6 µ in wall thickness, 7 -22 p. in diameter, 475 -968 µ in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma distinct, uniseriate; (b) paratracheal parenchyma sparse; few cells with tannin inclusions.

Rays nonstoried, heterogeneous, 1- 2- seriate, 2 -21 cells in height; almost all the cells with tannin inclusions.

10. Castanopsis kusanoi Hay. Mater. Fl. Formos. (1911) 302.

Syn. Castanopsis brevistella Hay. et Kaneh. ex A. Camus

Chat. Mong. Cast, et Castanop. (1929) 481, f. 16.

Castanopsis sinsuiensis Kaneh. Formos. Trees rev. ed.

(1936) 94. F. 49. Wood semi -diffuse porous; growth ring distinct, delineated by a band of thick -walled, flattened fibers. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 2 -14 per mm2 solitary, polygonal or orbicular, 3 -5 p. in wall thickness, 28 -193 µ in diameter; end walls oblique, perforation plates simple; pits 18 leading to contiguous tracheal cells bordered, pit cavity orbicular or elliptical, aperture lenticular (fusiform); vessel members 239 -648 p. in length. Tyloses present in heartwood. Vasicentric tracheids present, confined to the vicinity of larger vessels.

Fibers thin -walled, 2 -5 p, in wall thickness, 7 -24 p. in dia- meter, 598 -982 p. in length.

Parenchyma poorly differentiated against the background of fibrous tissue; (a) diffuse parencyma sparse; (b) apotracheal banded parenchyma abundant, in numerous, concentric, 1- 3- seriate, rag- ged tangential bands; (c) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath).

Rays nonstoried, heterogeneous, uniseriate, 1 -19 cells in height.

11. Castanopsis stellatospiria Hay. Mater. Fl. Formos. (1911) 302.

Syn. Castanopsis matsudai Hay. ex Camus Chat. Monog. Cast.

et Castanop. (1929) 482. f. 15.

Castanopsis brevispina Hay. Mater. Fl. Formos. (1911)

300.

Wood diffuse porous; growth ring distinct, delineated by a band of thick -walled fibrous tissue. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 6 -18 per mm2; 19 solitary, polygonal, orbicular or elliptical, 2 -4 µ in wall thickness,

59 -181 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbi- cular or elliptical, aperture lenticular; vessel members 139-512 µ in length. Tyloses present in heartwood. Vasicentric tracheids present, confined to the vicinity of large vessels.

Fibers 3 -6 µ in wall thickness, 8 -18 p. in diameter, 672 -995 µ in length. Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma abundant, conspicuous, in numerous, concentric, more or less continuous, 1- 3- seriate, tangential bands; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath), some cells with tannin inclusions.

Rays nonstoried, heterogeneous, uniseriate, 1 -25 cells in height, few cells with tannin inclusions.

12. Lithocarpus amygdalifolius (Skan) Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Syn. Quercus amygdalifolius Skan in Forbes et Hemsl. Ind.

Fl. Sin. II (1899) 506.

Pasania amygdalifolia (Skan) Schottky in Engl. Bot.

Jahrb. XLVII (1912) 188.

Wood diffuse porous; growth ring indistinct. 20

Pores unevenly distributed, arranged in flame -like or

stream -like clusters which extend for wome distance radially, 4 -19

per mm2; solitary, orbicular or oval; 3 -8 µ in wall thickness, 32-

168 µ in diameter; end walls oblique, perforation plates simple;

pits leading to contiguous tracheal cells bordered, ; pit cavity orbicu- lar, aperture lenticular; vessel members 141 -524,µ in length. Vas - icentric tracheids present, forming a 1- 2- seriate sheath, inter- rupted by parenchyma, about the larger vessels.

Fibers thick -walled, 3 -7 µ in wall thickness, 7 -22 µ in dia- meter, 432 -842 µ in length.

Parenchyma apotracheal banded or paratracheal; (a) banded parenchyma abundant, conspicuous, in numerous, concentric, con- tinuous, 1- 3- seriate (mostly 1- seriate) tangential bands; (b) para- tracheal parenchyma sparse, restricted to occasional cells, a few cells with tannin inclusions.

Rays nonstoried, heterogeneous; (a) aggregate rays com- posed of smaller rays separated by strands of fibrous tissue; (b) narrow rays numerous, uniseriate, 1 -19 cells in height; most of the cells with tannin inclusions.

13. Lithocarpus castanopsisifolius Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Syn. Quercus lepidocarpa Hay. Mater. F1. Formos. (1911) 291. 21

Pasania lepidocarpa (Hay.) Schottky in Engl. Bot. Jahrb.

XLVII (1912) 660.

Quercus castanopsisifolia Hay. Icon. Fl. Formos. III

(1913) 179, f. 26.

Pasania castanopsisifolia Hay. 1.. c.

Lithocarpus lepidocarpa Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Synaedrys amygdalifolia (Shan) Koidz. form, castanopsisi-

folia Kudo in Journ. Soc. Trap. Agr. Formos. III (1931)

386.

Synaedrys lepidocarpa (Hay.) Koidz. in Bot. Mag. Tokyo

XXX (1916) 189.

Wood diffuse porous; growth ring scarcely distinct, delin-

eated by a band of flattened fibers. Pores unevenly distributed, arranged in flame -like or stream-

like clusters ... which extend for some distance radially, 4 -16 per mm?

solitary, orbicular or elliptical, 2 -4 µ in wall thickness, 31 -176 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture orbicular or elliptical; vessel members 134 -586 µ in length. Tyloses present in heartwood. Vasicentric tracheids

present, confined to the vicinty of larger vessels.

Fibers 2 -6 µ in wall thickness, 7 -23 µ in diameter, 22

581 -987 µ in length. Parenchyma diffuse -in- aggregate and paratracheal; (a) dif- fuse-in-aggregate parenchyma conspicuous; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); a few cells with book - shaped crystals or tannin inclusions.

Rays ;nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 750 i in width, many cells (into the hundreds) and up to 5,1 21 µin height; aggregate rays composed of smaller rays separated by strands of fibrous tissue, otherwise comparable to the compound rays, few cells with crystals; (b) nar- row rays very numerous, unseriate, occasionally biseriate, 1 -23 cells in height.

14. Pasania kodaihoensis (Hay.) H. L. Li in Bull. Torrey Bot.

Club LXXX (1953) 321.

Syn. Lithocarpus kodaihoensis_. Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Quercus kodaihoensis Hay.Icon. Pl. Formos. IV (1914) 21.

Synaedrys kodaihoensis (Hay.) Kudo in Journ. Soc. Trop.

Agr. Formos. III (1931) 388.

Wood diffuse porous; growth ring indistinct. Pores unevenly distributed, arranged in flame -like or stream- 2 like clusters which extend for some distance radially, 3 -5 per mm2; solitary and occasionally in groups of 2, orbicular or elliptical, 23

3 -6 µ in wall thickness, 31 -196 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture lenticular; vessel members

149 -485 µ in length,. Tyloses present, numerous in heartwood. Vasicentric tracheids present, confined to the vicinity of larger vessels.

Fibers thick- walled, 3 -7 p. in wall thickness, 11 -34 µin diameter, 451 -896 p. in length. Parenchyma diffuse -in- aggregate and paratracheal; (a) dif- fuse-in- aggregate parenchyma abundant, conspicuous; (b) paratrach-

eal parenchyma sparse, restricted to occasional cells (not forming a sheath); few cells with book - shaped crystals.

Rays nonstoried, heterogeneous; (a) compound and aggregate

rays several- many- seriate, up to 165 p, in width, many cells (into hundreds) and up to 7,510 p. in height; aggregate rays composed of

smaller rays separated by strands of fibrous tissue, otherwise

comparable to the compound rays, few cells with book - shaped

crystals; very few sclereids present; (b) narrow rays very numerous,

uniseriate, occasionally biseriate, 1 -24 cells in height.

15. Pasania konishii (Hay.) Schottky in Engl. Bot. Jahrb. XLVII

(1912) 163.

Syn. Lithocarpus konishii Hay. Gen. Ind. Fl. Formos. (1916) 72 24

Quercus konishii Hay. Fl. Mont. Formos. (1908) 201,

t. 37.

Synaedrys konishii (Hay.) Koidz. in Bat. Mag. Tokyo

XXX (1916) 196.

Quercus cornea Lour. var. konishii Hay. Icon. Fl. For-

mos. III (1913) 179.

Wood diffuse porous; growth ring distinct, delineated by thick - walled fibers bordering on light colored early wood. Pores unevenly distributed, arranged in flame -like or stream -like clusters which extend for some distance radially.

15 -28 per mm2; solitary, orbicular or elliptical, 3 -7 µ in wall thick- ness, 28 -149 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture orbicular or lenticular; vessel members

144 -491 µ in length. Vasicentric tracheids present, forming a

1- 2- seriate sheath, interrupted by parenchyma, about the vessel.

Fibers 3 -6 p. in wall thickness, 7 -21 µ in diameter, 459-

1,060 p. in length.

Parenchyma conspicuous; (a) diffuse -in- aggregate parenchy- ma numerous; (b) apotracheal banded parenchyma in concentric, continuous, 1- 4- seriate, tangential bands; (c) paratracheal paren- chyma sparse, restricted to occasional cells; a few cells with book- shaped crystals. 25

Rays nonstoried, heterogeneous; (a) aggregate rays compos- ed of smaller rays separated by strands of fibrous tissue; (b) nar- row rays numerous, 1- 2- seriate, 1 -41 cells in height.

16. Pasania brevicaudata (Skan) Schottky in Engl. Bot. Jahrb.

XLVII (1912) 666.

Syn. Lithocarpus brevicaudata (Skan) Hay. Gen. Ind. Fl.

Formos. (1916).

Quercus brevicaudata Skan in Journ. Linn. Soc. XXVI

(1899) 308.

Quercus impressivena Hay. Mater. Fl. Formos. (1911)

291.

Pasania impressivena (Hay. ) Schottky in 1. c. 675.

Synaedrys brevicaudata (Skan) Koidz. in Bot. Mag. Tokyo

XXX (1916) 194.

Lithocarpus impressivena (Hay.) Hay. Gen. Ind. Fl,

Formos., (1917) 72.

Synaedrys impressivena Masam. in Aim. Rep. Taihoku

Bot. Gard. II (1932) 14.

Synaedrys brevicaudata var. pinnativena Suzuki ex

Masam. Short Fl. Formos. (1936) 43,, Lithocarpus brevicaudata var. pinnativena Yamamoto

in Journ. Soc. Trop. Agr. Formos. IV (1932) 49. 26

Wood diffuse porous; growth ring scarcely distinct, delinea- ted by a faint narrow line of denser fibrous tissue at the outer margin. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 6 -25 per mm2 solitary, orbicular or elliptical, 3 -5 µ in wall thickness, 25 -189 µ in diameter; end walls oblique, perforation plates simple; pits lead- ing to contiguous tracheal cells bordered, pit cavity orbicular, ap- erture lenticular; vessel members 132 -558 p. in length. Tyloses present in heartwood. Vasicentric tracheids present, forming a

1- 2- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers thin -walled, 2 -5 µ in wall thickness, 6 -26 p, in diam- eter, 498 -972 µ in length. Parenchyma apotracheal diffuse, banded and paratracheal; (a) diffuse parenchyma sparse, scattered among fibers as single cells; (b) banded parenchyma abundant, conspicuous, in numerous, concentric, continuous, 1- 3- seriate, tangential bands; (c) paratra- cheal parenchyma sparse, restricted to occasional cells. Rays nonstoried, heterogeneous (marginal cells upright);

(a) compound and aggregate rays several- many- seriate, up to

152 µ in width; aggregate rays composed of smaller rays separated by strands of fibrous tissue, otherwise comparable to the compound rays; (b) narrow rays numerous, uniseriate, 1 -16 cells in height; many cells with tannin inclusions. 27

17. Pasania uraiana (Hay.) Schottky in Engl. Bot. Jahrb. XLVII

(191 2) 675.

Syn. Quercus uraiana Hay. Mater. F1, Formos. (1911) 299.

Lithocarpus uraiana Hay. Gen. Ind. Fl. Formos. (1916) 72,

Synaedrys uraiana Koidz. in Bot. Mag. Tokyo XXX (1916)

198.

Shiia uraiana Kaneh. et Hatus. Formos, Trees rev, ed.

(1936) 116. in notes.

Castanopsis uraiana (Hay. ) Kaneh. et Hatus, in Trans.

Nat. Hist. Soc. Formos. XXIX (1939) 155. Limlia uraiana Masam. et Tomiya in Acta Bot. Taiwan

I(1947) 1.

Wood ring porous; growth ring conspicuous, delineated by thick -walled fibrous tissue bordering on early wood. Pores unevenly distributed, early wood pores very large forming a narrow, conspicuous band 1 -3 pores in width, 2 -9 per mm2; solitary, polygonal, orbicular or elliptical, 3 -5 p. in wall thickness, 52 -183 µ in diameter; late wood pores small, polygonal, arranged in flame -like patches. 69 -149 per mm2, the transition in pore size from early to late wood abrupt; end walls oblique, per- foration plates simple or those in the smaller late wood vessels oc- casionally scalariform, 1 -4 bars; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture orbicular or lenticular; 28

vessel members 230 -634 µ in length. Tyloses present, abundant in heartwood. Vasicentric tracheids present, confined to the vicinity

of the early wood vessels.

Fibers 3 -7 µ in wall thickness, 7 -21 µ in diameter, 383 -996 p. in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma abundant, in numerous, concentric, continuous, 1-3-

seriate, tangential bands; (b) paratracheal parenchyma sparse, re-

stricted to occasional cells.

Rays nonstoried, heterogenous; (a) aggregate rays composed

of smaller rays separated by strands of fibrous tissue; (b) narrow

rays very numerous, uniseriate, 1 -21 cells in height.

18. Pasania kawakamii (Hay. ) Schottky in Engl. Bot. Jahrb. XLVII

(1912) 666.

Syn. Lithocarpus kawakamii Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Quercus kawakamii Hay. Fl. Mont. Formos. (1908) 201.

Synaedrys kawakamii (Hay.) Koidz, in Bot. Mag. Tokyo

XXX (1916) 196.

Wood diffuse porous; growth ring distinct. Pores unevenly distributed, arranged in flame -like or

stream -like clusters which extend for some distance radially, 2 -12 29 per mm2; solitary, orbicular or elliptical, 3 -5 p, in wall thickness,

49 -189 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicu- lar, aperture orbicular or lenticular; vessel members 195 -510 µ in length. Tyloses present in heartwood. Vasicentric tracheids pres- ent, confined to the vicinity of larger vessels.

Fibers 2 -5 µ in wall thickness, 8 -25 p. in diameter, 501 -976 µ in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma very abundant, conspicuous, in numerous, concentric, continuous, 1- 4- seriate (mostly 2- 3- seriate), tangential bands;

(b) paratracheal parenchyma sparse, restricted to occasional cells

(not forming a sheath); some cells with tannin inclusions.

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 159 µ in width, many cells (into the hundreds) up to 13,920 µ in height; aggregate rays composed of smaller rays separated by strands of fibrous tissue, otherwise com- parable to the compound rays; (b) narrow rays very numerous, uni- seriate, occasionally biseriate, 1 -27 cells in height; most of the cells with tannin inclusions.

19. Pasania dodoniaefolia Hay. Icon. Pl, Formos. III (1913) 181,

t, 27. 30

Syn. Lithocarpus dodoniaefolia Hay. Gen. Ind. F1. Formos.

(1916) 72.

Quercus dodoniaefolia Hay. Icon. Pl. Formos. III (1913)

181, t. 27.

Synaedrys formosana (Skan) Koidz. var. dodoniaefolia

Kudo in Journ. Soc. Trop. Agr. Formos. III (1931) 387.

Wood diffuse porous; growth ring distinct. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 5 -19 per mm? solitary, orbicular or elliptical, 3 -5 µ. in wall thickness, 42 -168 µ in diameter; end walls oblique, perforation plates simple; pits lead- ing to contiguous tracheal cells bordered, pit cavity orbicular, aper- ture lenticular; vessel members 179-691 µ in length. Tyloses pres- ent in heartwood. Vasicentric tracheids present, confined to the vicinity of larger vessels.

Fibers 2 -6 I. in wall thickness, 7 -18 µ in diameter, 546 -895 µ in length.

Parenchyma apotracheal banded and paratracheal; (a) band- ed parenchyma very abundant, conspicuous, in numerous, concen- tric, continuous, 1 -3 (mostly 2 -3) seriate bands; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); some cells with tannin inclusions.

Rays nonstoried, heterogeneous; (a) compound and aggregate 31

rays several- many- seriate, up to 225 µ in width, many cells in height; aggregate rays composed of smaller rays separated by

strands of fibrous tissue, otherwise comparable to the compound

rays; (b) narrow rays numerous, uniseriate, 1 -31 cells in height;

most of the cells with tannin inclusions.

20. Pasania randaiensis (Hay.) Schottky in Engl. Bot. Jahrb.

XLVII (191 2) 672.

Syn. Lithocarpus randaiensis Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Quercus randaiensis Hay. Mater. Fl. Formos. (1911) 295.

Synaedrys randaiensis Koidz. in Bot. Mag. Tokyo XXX

(1916) 197,

Lithocarpus rhombocarpa Hay. Gen. Ind. Fl, Forraos.

(1916) 72.

Quercus rhombocarpa Hay. Icon. Pl. Formos. III (1913)

186.

Pasania rhombocarpa Hay. 1, c.

Lithocarpus buisanensis Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Lithocarpus nakaii Hay. Icon. Fl. Formos. IX (1920)

106, f. 34.

Lithocarpus suisha.e.nsis Kaneh. et Yamam, in Yamam, Suppl. Icon, Pl. Formos. III (1927) 11, f. 4. 32

Synaedrys rhombocarpa (Hay.) Kudo in Journ. Soc. Trop.

Agr. Formos. III (1931) 388.

Synaedrys nakaii (Hay.) Kudo in 1. c.

Synaedrys rhombocarpa (Hay.) kudo form. suishaensis

Kudo in 1. c. 389.

Pasania suishaensis Nak. in Journ. Jap. Bot. XV

(1932) 275.

Pasania nakaii (Hay.) Nak. in Journ. Jap. Bot. XV

(1932) 272

Wood diffuse porous; growth ring scarcely distinct.

Pores unevenly distributed, arranged .. in flame -like or stream -like clusters which extend for some distance radially,

11 -31 per mm2; solitary, orbicular or elliptical, 3 -5 µ in wall thickness, 30 -189 p. in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture orbicular or lenticular; vessel members 86 -432 p. in length. Tyloses present in heartwood.

Vasicentric tracheids present, forming a 1- 3- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers 3 -6 µ in wall thickness, 7 -23 p. in diameter, 452-

963 µ in length. 33

Parenchyma abundant, conspicuous, distributed in (a) dif-

fuse-in- aggregate or (b) irregularly apotracheal banded; (c) para-

tracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); a few cells with tannin inclusions.

Rays nonstoried, heterogeneous; (a) Broad rays aggregate,

composed of smaller rays separated by strands of fibrous tissue,

many cells (into the hundreds) and up to 13,490 µin height; (b) nar-

row rays numerous, uniseriate, 1 -32 cells in height; a few cells with tannin inclusions.

21. Pasania formosana (Skan) Schottky in Engl. Bot. Jahrb. XLVII

(1912) 670.

Syn. Lithocarpus formosana (Skan) Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Quercus formosana Skan in Journ. Soc. Bot. XXVI

(1899) 513.

Synaedrys formosana (Skan) Koidz. in Bot. Mag. Tokyo

XX (1916) 195.

Wood diffuse porous; growth ring indistinct.

Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 3 -13 per mm2;2

solitary and occasionally in groups of 2, orbicular or elliptical, 34

3 -4 µ in wall thickness, 49 -133 µ in diameter; end walls oblique,

perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture lenticular,; vessel members

179 -573 H. in length. Tyloses present in heartwood. Vasicentric

tracheids present, confined to the vicinity of larger vessels.

Fibers thick -walled, 3 -7 µ in wall thickness, 7 -18 µ in diam-

eter, 381 -863 µ in length.

Paranchyma diffuse -in- aggregate and paratracheal; (a) dif- fuse-in- aggregate parenchyma abundant; (b) paratracheal paren- chyma sparse, restricted to occasional cells (not forming a sheath);

some cells with tannin inclusions, a few cells with book - shaped crystals. Rays nonstoried, heterogeneous, (a)compound and aggregate

rays several- many- seriate, up to 238 µ in width, many cells (into the hundreds) and up to 1,028 p. in height; aggregate rays composed

of smaller rays separated by strands of fibrous tissue, otherwise comparable to the broad rays; (b) narrow rays very numerous, uniseriate, 1 -31 cells in height; most of the cells with tannin in-

clusions; few cells with book - shaped or polyhedral crystals.

22. Pasania ternaticupula (Hay.) Schottky in Engl. Bot. Jahrb.

XLVII (1912) 675.

Syn. Quercus ternaticupula Hay.Mater. Fl. Formos. (1911) 298. 35

Quercus subreticulata Hay. Icon. Pl. Formos. III (1913)

184.

Lithocarpus subreticulata Hay. Gen. Ind. F1. Formos.

(1916) 72. Synaedrys ternaticupula (Hay.) Koidz. in Bot. Mag. Tokyo

XXX (1916) 198.

Lithocarpus ternaticupula Hay. Gen. Ind. Fl. Formos.

(1916) 72.

Lithocarpus matsudai Hay. Icon. Pl. Formos. IX (1920)

108, f. 35. Cyclobalanopsis ternaticupula (Hay.) Kudo in Journ. Soc.

Trop. Agr. Formos. III (1931).390.

Synaedrys matsudai (Hay.) Kudo in 1. c. 388. Lithocarpus ternaticupula Hay. var. sinsuiensis Kaneh.

Formos. Trees rev, ed. (1936) 116, f. 36. Synaedrys kuarunensis Tomiya in Trans. Nat. Hist. Soc.

Formos. XXXVI (1942) 346.

Wood diffuse porous; growth ring distinct, delineated by

denser fibers bordering on early wood. Pores unevenly distributed, arranged in flame -like or stream- like clusters (sometimes in broader patches) which extend for some

distance radially, 10 -51 per mm2; solitary, orbicular or elliptical,

4 -7 µ in wall thickness, 30 -173 µ in diamter; end walls oblique, 36 perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture lenticular; vessel members

81 -508 µ.in length. Tyloses occasionally present in heartwood. Vas- icentric tracheids present, forming a 1- 3- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers 3 -7 p. in wall thickness, 7 -20 µ in diameter, 392 -804 µ in length. Parenchyma abundant, conspicuous, apotracheal banded and paratracheal; (a) banded (sometimes diffuse -in- aggregate) paren- chyma in concentric, more or less continuous, 1- 3- seriate ragged bands; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); many cells with tannin inclusions.

Rays nonstoried, heterogeneous, (a) compound and aggregate

rays several -many- seriate, up to 163 µ in width, many cells (into

the hundreds) and up to 6, 213 µ in height; many sclereids and book - shaped crystals in heartwood rays; aggregate rays composed of

smaller rays separated by strands of fibrous tissue; (b) narrow rays numerous, uniseriate, 1 -18 cells in height; many cells with tannin inclusions.

23. Pasania nantoensis (Hay.) Schottky in Engl. Bot. Jahrb. XIVII

(1912) 675.

Syn. Quercus nantoensis Hay. Mater. Fl. Formos. (1911) 293. 37

Lithocarpus nantoensis Hay. Gen. Ind. Fl, Formos.

(1916) 72.

Synaedrys nantoensis (Hay.) Koidz. in Bot. Mag. Tokyo

XXX (1916) 199.

Wood diffuse porous; growth ring indistinct.

Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 10 -25 per 2 mm solitary, orbicular or elliptical, 3 -7 µ in wall thickness, 29-

159 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbic- ular, aperture orbicular or lenticular; vessel members 142 -528 µ in length. Tyloses present in heartwood. Vasicentric tracheids present, forming a 1- 3- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers 3 -7 µ in wall thickness, 7 -20 p. in diameter, 368 -896

µ in length. Parenchyma abundant, conspicuous, apotracheal banded and paratracheal; (a) banded parenchyma in numerous, concentric, more or less continuous, 1- 2- seriate, tangential bands; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); a few cells with tannin inclusions.

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 309 11 in width, many cells (into 38

the hundreds) and up to 7,123 µ in height; aggregate rays composed of

smaller rays separated by strands of fibrous tissue, otherwise com-

parable to the compound rays; (b) narrow rays numerous, uniseriate,

1 -22 cells in height; a few cells with tannin inclusions.

24. Quercus variabilis Bl. in Mus. Bot. Lugd -Bot. I (1850) 297.

Syn. Quercus serrata sensu Carruth. in Journ. Linn. Soc.

VI (1862) 32.non Thunb.

Quercus chinensis sensu Bunge in Mem. Say. Etr. Acad.

Sci. St. Petersberg II (1835) Enum. Pl. Chin. Bor.

61; non. Abel.

Quercus bungeana Forbes in Journ. Bot. XXII (1884) 83. Wood semi -diffuse porous; growth ring distinct, delineated through an abrupt difference in size between the pores of the late wood and those in the early wood of the succeeding ring.

Pores scattered, those in the early wood large, decreasing gradually in size toward the outer margin of the ring, 29 -53 per mm2; solitary, orbicular or elliptical, 3 -7 H. in wall thickness,

29 -219 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbic- ular or elliptical, aperture lenticular; vessel members 115 -384 µ in length. Tyloses present in heartwood. Vasicentric tracheids present, forming a 1- 3- seriate sheath, interrupted by parenchyma, about the vessels. 39

Fibers thick -walled, abundant in late wood, sometimes

gelatinous; 3 -7 µ in wall thickness, 7 -18 µ in diameter, 528 -893 p. in length.

Parenchyma conspicuous; (a) diffuse -in- aggregate paren-

chyma abundant, distributed at random; (b) apotracheal banded

parenchyma in concentric, continuous, 1- 3- seriate ragged tangential

bands; (c) paratracheal parenchyma sparse, restricted to occasional

cells (not forming a sheath); a few cells with book - shaped crystals or tannin inclusions.

Rays nonstoried, homogeneous; (a) compound and aggregate

rays several- many- seriate, up to 719 µin width, many cells (into the hundreds) and up to 4,192 pin height; aggregate rays composed

of smaller rays separated by strands of fibrous tissue; (b) narrow

rays numerous, uniseriate, 1 -27 cells in height; many cells with book- shaped crystals or tannin inclusions.

25. Quercus acutissima Carruth. in Journ. Linn, Soc. VI (162) 33.

Syn. Quercus serrata sensu Sieb. et Zucc. in Abh. Akad.

Muench. IV 3. (1846) 226. non Thunb.

Wood semi -diffuse porous; growth ring distinct, delineated through an abrupt difference in size between the pores of the late wood and those in the early wood of the succeeding ring.

Pores scattered, those in the early wood large, decreasing 40

gradually in size toward the outer margin of the ring, 10 -41 per 2 mm ; solitary and occasionally in radial groups of 2; orbicular or

elliptical, 3 -5 p, in wall thickness, 29 -159 µ in diameter; end walls

oblique, perforation plates simple; pits leading to contiguous tra- cheal cells bordered, pit cavity orbicular or elliptical, aperture

orbicular or lenticular; vessel members 141 -431 µ in length. Few tyloses present in heartwood. Vasicentric tracheids present, form- ing a 1- 4- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers thick -walled, sometimes gelatinous, 3 -7 p in wall thickness, 8 -19 µ in diameter, 489 -963 p. in length.

Parenchyma conspicuous; (a) diffuse -in- aggregate paren- chyma sparse, restricted to occasional cells (not forming a sheath); a few cells with book - shaped crystals or tannin inclusions. Rays nonstoried, homogeneous (very few heterogeneous) or nearly so; (a) compound and aggregate rays several- many- seriate, up to 628 i in width, many cells (into the hundreds) and up to 5,123 µ in height; aggregate rays sparse; (b) narrow rays very numerous, uniseriate, 1 -28 cells in height; a few cells with book - shaped crys- tal or tannin inclusions.

26. Quercus phillyraeoides A. Gray, Mem. Am. Acad. N. S. VI

(1858 -59) 406. 41

Syn. Quercus Ilex L. var. phillyraeoides Franch. in Journ. de

Bot. (1899) 152.

Quercus Ilex L. var. acordonda Skan in Journ. Linn. Soc.

XXVI (1899) 516.

Quercus acrodonta Seem. in Engl. Bot. Jahrb. XXIII

No. 57 (1897) 48; XXIX (1900) 290,

Quercus phillyraeoides (Sp. name orig. philliraeoides) Gray var. sinensis Schottky in Engl. Bot. Jahrb. XLVII

(1912) 643.

Wood diffuse porous; growth ring scarcely distinct.

Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 3 -48 per mm2; solitary, orbicular or elliptical, 3 -7 µ in wall thickness, 193 -522 p. in diameter; end walls oblique, perforation plates simple; pits lead- ing to contiguous tracheal cells bordered, pit cavity orbicular, ap- erture orbicular or lenticular; vessel members 127 -409 p. in length. Tyloses present in heartwood. Vasicentric tracheids present, sparse, sometimes forming a uniseriate sheath, interrupted by parenchyma, about the larger vessels.

Fibers thick -walled, sometimes gelatinous, 3 -7 p. in wall thickness, 7 -19 p. in diameter, 497 -942 p. in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma abundant, conspicuous, in numerous, concentric, 42 continuous, 1- 3- seriate (mostly 1 -2) ragged tangential bands;

(b) paratracheal parenchyma very sparse; few cells with book- shaped or polyhedral crystals.

Rays nonstoried, homogeneous; (a) compound and aggregate rays several- many- seriate, up to 526 µ in width, many cells (into the hundreds) and up to 12,750 p. in height; aggregate rays very sparse; (b) narrow rays very numerous, uniseriate, occasionally biseriate, 1 -19 cells in height; few cells with book - shaped crystals or tannin inclusions; sclereids occasionally present in groups.

27. Quercus semicarpifolia Smith Subsp. glabra (Franch.) Handel -

Mazzetti in Symb. Sin. VII (1929) 39, Syn. Quercus Ilex L. var. spinosa sensu Hay. Mater. Fl.

Formos. (1911) 290. non Franch.

Quercus spinosa A. David var. miyabei Hay. Icon, Pl.

Formos, VII (1918) 37.

Quercus tatakaensis Tomiya in Trans. Nat. Hist. Soc.

Formos. XXXIV (1942) 346.

Wood diffuse porous; growth ring scarcely distinct. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 4 -47 per mm2; solitary, orbicular or elliptical, 3 -6 µ in wall thickness, 31 -116 µ in diameter; end walls oblique, perforation plates simple; pits 43 leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture lenticular; vessel members_ 74:- 3O2;µ in length. Tyloses present in heartwood. Vasicentric tracheids present, forming a uniseriate sheath, interrupted by parenchyma, about the vessels.

Fibers 3 -6 µ in wall thickness, 7 -22 µ in diameter, 397 -9O8µ in length; many gelatinous fibers in early wood. Parenchyma diffuse -in- aggregate and paratracheal; (a) dif- fuse-in- aggregate parenchyma abundant, conspicuous; (b) para- tracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); few cells with tannin inclusions.

Rays nonstoried, homogeneous or with very few upright mar- ginal cells; (a) broad rays mostly aggregate which are composed of smaller rays separated by strands of fibrous tissue; (b) narrow rays very numerous, uniseriate, 1 -23 cells in height; most of the cells with tannin inclusions; few cells with book- shaped or polyhedral crystals.

28. Cyclobalanopsis pachyloma (O. Seem.) Schottky in Engl. Bot.

Jahrb. XLVII (1912) 650.

Syn. Quercus pachyloma O. Seem, in Engl. Bot. Jahrb. Beibl.

XIII (1897) 54.

Quercus tomentosicupula Hay. Icon. Pl. Formos. III

(1913) 185, f. 33. 44

Cyclobalanopsis tomentosicupula Hay. 1. c. Cyclobalanopsis pachyloma Schottky var. yamamotoi Kudo

in Journ. Soc. Trop. Agr. Formos. II (1930) 148. syn.

nov.

Quercus pachyloma O. Seem. var. tomentosicupula (Hay.)

Metcalf. Fl. Fukien I (1942) 76.

Wood diffuse porous; growth ring indistinct. Pores unevenly distributed, arranged in flame -like or stream- 2 like clusters which extend for some distance radially, 7 -22 per mm,; solitary, orbicular or elliptical, 3 -6 µ in wall thickness, 30 -188 µ in diameter; end walls oblique, perforation plates simple; pits lead- ing to contiguous tracheal cells bordered, pit cavity orbicular, aperture orbicular or lenticular; vessel members 159 -542 µ.inlength.

Vasicentric tracheids present, 1- several- seriate, sometimes form- ing large, confluent groups about the pore clusters.

Fibers thick -walled, sometimes gelatinous, 3 -7 µ in wall thickness, 7 -22 µ in diameter, 542-1,039 µ in length. Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma conspicuous, in numerous, concentric, continuous, 1 -2- seriate, ragged tangential bands; (b) paratracheal parenchyma sparse, restricted to occasional cells; very few cells with crystals; a few cells with tannin inclusions.

Rays nonstoried, heterogeneous; (a) compound and aggregate 45 rays several- many- seriate, up to 312 µ in width, many cells (into the hundreds) and up to 5, 863 µ in height; (b) narrow rays numerous, uniseriate, 1 -21 cells in height, a few cells with tannin inclusions.

29. Cyclobalanopsis morii (Hay.) Schottky in Engl. Bot. Jahrb.

XLVII (191 2) 658.

Syn. Quercus morii Hay. Mater. Fl. Formos. (1911) 292.

Wgod diffuse porous, growth ring distinct, delineated by denser fibers bordering on early wood. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 3 -15 per mm? solitary (very few in groups of 2 -3), polygonal, elliptical or oval,

3 -5 µ in wall thickness, 35 -196 p. in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture lenticular; vessel members

135 -501 ti, in length. Vasicentric tracheids present, forming a uni- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers thin -walled, sometimes gelatinous, 2 -4 p. in wall thickness, 10 -32 µ in diameter, 432 -961 µ in length.

Parenchyma diffuse -in- aggregate and paratracheal; (a) dif- fuse-in- aggregate parenchyma abundant, sometimes in large clus- ters; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); very few cells with crystals. 46

Rays nonstoried, heterogeneous; (a) aggregate rays composed

of smaller rays separated by strands of fibrous tissue; (b) narrow

rays numerous, uniseriate, occasionally biseriate, 1 -55 cells in height; few cells with tannin inclusions.

30. Cyclobalanopsis championii (Benth.) Oerst. in Engl. Bot. Jahrb.

XLVII (1912) 652, et Kjoeb. Vidensk. Meddel. (1866) 79.

Syn. Quercus championii Benth. in Hook. Journ. Bot. VI (1854)

113.

Wood diffuse porous; growth ring indistinct. Pores unevently distributed, arranged in flame -like or stream -like clusters which extend for some distance radially, 2 -20 per mm2; solitary, orbicular or elliptical, 3 -5 µ in wall thickness,

33 -152 µ in diameter; end walls oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbic- ular, aperture lenticular; vessel members 119 -486 µ in length.

Vasicentric tracheids present, forming a 1- 3- seriate sheath, inter- rupted by parenchyma, about the vessels.

Fibers thick -walled, sometimes gelatinous, 3 -6 µ in wall thickness, 7 -27 µ in diameter, 532 -967 µ in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parencyma abundant, conspicuous, in numerous, concentric, con- tinuous, 1- 3- seriate ragged tangential bands; (b) paratracheal 47

parenchyma sparse, restricted to occasional cells (not forming a

sheath); many cells with book - shaped crystals or tannin inclusions.

Rays nonstoried, heterogeneous; (a) compound rays several -

many- seriate, up to 273 µ in width, many cells (into the hundreds)

and up to 10,500 µ in height; aggregate rays sparse; (b) narrow rays

very numerous, uniseriate, occasionally biseriate, 1 -24 cells in height; a few cells with book- shaped crystals or tannin inclusions.

31. Cyclobalanopsis gilva (Blume) Oerst. in Kjoeb. Vidensk. Meddel.

XVIII (1866) 78.

Syn. Quercus gilva .Blume in Mus. Bot. Lugd. Bot. I (1850) 306.

Wood diffuse porous; growth ring distinct, delineated by a

narrow line of fibrous tissue at the outer margin.

Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 3 -18 per mm?

solitary, orbicular, 3 -5 µ in wall thickness, 28 -47 p. in diameter;

end walls oblique, perforation plates simple; pits leading to contig- uous tracheal cells bordered, pit cavity orbicular, aperture lenti- cular; vessel members 134 -452 µ in length. Vasicentric tracheids present, forming a 1- 3- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers thin- walled, occasionally gelatinous, 2 -5 µ in wall thickness, 7 -18 µ in diameter, 568' -761 11 in length. 48 Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma abundant, conspicuous, in numerous, concentric, con- tinuous, 1- 3- seriate, ragged tangential bands; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); very few cells with book - shaped crystals.

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 345 µ in width, many cells (into the hundreds) and up to 2,510 p. in height, aggregate rays composed of smaller rays separated by strands of fibrous tissue, otherwise comparable to the compound rays; (b) narrow rays very numerous, uniseriate, 1 -20 cells in height; few cells with tannin inclusions; very few cells with book - shaped crystals.

32. Cyclobalanopsis stenophylloides (Hay.) Kudo et Masam. in

Trans. Nat. Hist. Soc. Formos. (1930) 149.

Syn. Quercus stenophylloides Hay. Icon. Pl. Formos. IV

(1914) 21.

Wood diffuse porous, growth ring scarcely distinct. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 2 -17 per mm2 solitary, polygonal or orbicular, 2 -5 µ in wall thickness, 38 -154 µ in diameter; end walls oblique, perforation plates simple; pits lead- ing to contiguous tracheal cells bordered, pit cavity orbicular, 49 aperture lenticular; vessel members 119 -517 µ in length. Tyloses present in heartwood. Vasicentric tracheids present, forming a

1- 3- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers sometimes gelatinous, 3 -6 µ in wall thickness, 7 -21

µ in diameter, 539 -967 µ in length. Parenchyma diffuse, diffuse -in- aggregate and paratracheal;

(a) diffuse parenchymaubiquitous; (b) diffuse -in- aggregate paren- chyma conspicuous, sometimes in more or less continuous, 1 -2- seriate ragged tangential bands; (c) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); many cells with book - shaped crystals.

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 375 p. in width, many cells (into the hundreds) and up to 5,983 p. in height, a few cells with book - shaped crystals; aggregate rays composed of smaller rays separated by strands of fibrous tissue, otherwise comparable to the compound rays; (b) narrow rays very numerous, uniseriate, 1 -26 cells in height.

33. Cyclobalanopsis glauca (Thunb.) Oerst. in Kjoeb. Vidensk.

Meddel. XVIII (1866) 70.

Syn. Quercus glauca Thunb. Fl. Jap. (1784) 175.

Quercus annulata Smith in Raes. Cyclop.. XXIX No..22 (1819). 50

Quercus phullata Hamilton apud D. Don Prodr. Fl. Nepal

(1825) 57.

Perytis glauca Rafin Alsog. Am. (1838) 29.

Quercus glauca Thunb. var. caesia Blume Mus. Bot, Ludg.

Bot. I (1850) 303.

Quercus sasakii Kaneh. ex Hay. Icon. Pl. Formos, VI

(1916) 64.

Quercus laxiflora Lindley in Wallich Cat. No. 2774

(nomen nudum) (1929).

Quercus dentosa Lindley in 1. c, 2775. Cyclobalanopsis sasakii (Kaneh.) Kudo et Masan), in Trans.

Nat. Hist. Soc. Formos, XX (1930) 161.

Wood diffuse porous; growth ring scarcely distinct. Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 3 -14 per mm2 solitary, orbicular or elliptical, 4 -7 µ in wall thickness, 40 -168 µ in diameter; end walls slightly oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbic- ular, aperture lenticular; vessel members 109 -441 µ in length. Tyloses present, sparse, in heartwood. Vasicentric tracheids present, forming a 1- 3- seriate sheath, interrupted by parenchyma, about the vessels.

Fibers sometime gelatinous, thick -walled, 3 -7 µ in wall 51 thickness, 8 -22 µ in diameter, 462 -907 µ in length. Parenchyma diffuse -in- aggregate and paratracheal; (a) dif- fuse-in- aggregate parenchyma numerous, conspicuous, many larger cells with book - shaped crystals; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath).

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 405 µ in width, many cells (into the hundreds) and up to 3,374 p. in height, most of the cells with book - shaped crystals; many highly sclerified cells present in heart- wood; aggregate rays sparse; (b) narrow rays numerous, uniseriate, occasionally biseriate, 1 -22 cells in height; most of the cells with tannin inclusions.

34. Cyclobalanopsis longinux (Hay. ) Schottky in Engl. Bot. Jahrb.

XLVII (1812) 657.

Syn. Quercus longinux Hay. Mater. Fl. Formos. (1911) 292. Quercus pseudomyrsinaefolia Hay. Mater. Fl. Formos.

(1911) 292.

Quercus taichuensis Hay. 1. c. 296. Cyclobalanopsis pseudomyrsinaefolia Hay. Mater. Fl.

Formos. (1911) 292.

Cyclobalanopsis taichuensis (Hay.) Schottky in 1. c. 683.

Cyclobalanopsis kaneherai Nak. In Journ. Jap. Bot. XV (19.39) 198. 52

Wood diffuse porous; growth ring indistinct.

Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 1 -10 per mm2 solitary, orbicular or elliptical, 3 -5 µ in wall thickness, 59 -250 p in diameter; end walls slightly oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbic- ular, aperture lenticular; vessel members 121 -512 µ in length. Ty- loses present in heartwood. Vasicentric tracheids present, forming a 1- 3- seriate sheath, interrupted by parenchyma, about the vessels, sometimes forming confluent, large patches about the pore clusters.

Fibers thick -walled, very few gelatinous, 3 -7 p in wall thick- ness, 8 -25 µ in diameter, 501 -993 µ in length.

Parenchyma apotracheal banded and paratracheal; (a) banded parenchyma conspicuous, in numerous, concentric, continuous,

1- 3- seriate, ragged tangential bands; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath); few cells with tannin inclusions.

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 453 µ in width, many cells (into the hundreds) and up to 5, 251 µ in height, a few cells with book - shaped crystals; aggregate rays sparse; (b) narrow rays numerous, uniseriate, occasionally biseriate, 1 -29 cells in height, most of the cells with tannin inclusions. 53

35. Cyclobalanopsis globosa Lin et Liu in Bull. Taiwan For. Res.

Inst. 110 (1965).

Wood diffus porous; growth ring indistinct.

Pores unevenly distributed, arranged in flame -like or stream- like clusters which extend for some distance radially, 4 -11 per mm2; solitary, elliptical, 3 -5 µ, in wall thickness, 41 -203 µ in diameter; end walls slightly oblique, perforation plates simple; pits leading to contiguous tracheal cells bordered, pit cavity orbicular, aperture lenticular; vessel members 119 -451 p. in length. Very few tyloses present in heartwood. Vasicentric tracheids present, forming a

1- 3- seriate sheath; interrupted by parenchyma, about the vessels.

Fibers thick -walled, 3 -7 µin wall thickness, 7 -19 p. in diam- eter, 503 -992 µ in length. Parenchyma diffuse -in- aggregate and paratracheal; (a) dif- fuse-in- aggregate parenchyma ubiquitous, conspicuous, sometimes reticulate, a few cells with book - shaped crystals; (b) paratracheal parenchyma sparse, restricted to occasional cells (not forming a sheath).

Rays nonstoried, heterogeneous; (a) compound and aggregate rays several- many- seriate, up to 495 µ in width, many cells (into the hundreds) and up to 3,724 µ, in height, a few cells with book - shaped crystals; aggregate rays sparse; (b) narrow rays very numerous, uniseriate, occasionally biseriate, 1 -32 cells in height, a few cells 54 with tannin inclusions. 55

ANATOMICAL KEY TO SPECIES OF FAGACEAE IN TAIWAN

1. Vasicentric tracheids absent Fagus hayatae

1. Vasicentric tracheids present 2

2. Gelatinous fibers absent 3

2. Gelatinous fibers present 24

3. Rays homogeneous 4

3. Rays heterogeneous 5

4. Scalariform perforations in late wood Castanea crenata

4. Porous perforations in late wood Castanea mollissirra

5. Pores polygonal 6

5. Pores not polygonal 14

6. Wood clearly ring porous 7

6. Wood semidiffuse or diffuse porous 10

7. Scalariform perforations in late wood 8

7. Porous perforations in late wood 9

8. Fibers 3 -7 p, in wall thickness Pasania uraiana

8. Fibers 2 -5 µ in wall thickness Castanopsis carlesii var. sessilis

9. Some rays compound or aggregate Castanopsis hystrix

9. All rays narrow Castanopsis kawa -_. kamii

10. Wood semidiffuse porous Castanopis kusanoi

10. Wood diffuse porous 11 56

11. Only porous performations in late wood 12

11. Some scalariform perforations in late Castanopsis carlesii wood

12. Parenchyma diffuse -in- aggregate and Castanopsis sub - paratracheal acuminata

12. Parenchyma apotracheal banded and paratracheal 13

13. Parenchyma crystals present, tannins Castanopsis for -

absent ` inosàna

13. Parenchyma crystals absent, tannins Castanopsis stella- present tospina

14. Growth ring indistinct or scarcely distinct 15

14. Growth ring distinct 19

15. Growth ring scarcely distinct Lithocarpus castan- opsisfolius

15. Growth ring indistinct 16

16. Parenchyma apotracheal banded and paratracheal 17

16. Parenchyma diffuse -in- aggregate and paratracheal 18

17. Tyloses absent in heartwood Lithocarpus amyg- dalìfolius

17. Tyloses present Pasania nantoensis

18, Tannin inclusions absent Pasania kodainoensis

18. Tanin inclusions present in parenchyma Pasania formosana

19. Tyloses absent Pasania konishii

19. Tyloses present in heartwood 20 57

20. Parenchyma apotracheal banded and paratracheal 21

20. Parenchyma diffuse, banded and paratracheal or diffuse -in- aggregate, banded and paratracheal 23

21. Crystals present in ray cells Pasania ternaticupu- la

21. Crystals absent 22

22. Pit aperture orbicular and lenticular Pasania kawakamii

22. Pit aperture lenticular only Pasania dodoniaefolia

23. Parenchyma diffuse banded and paratracheal Pasania brevicaudata

23. Parenchyma diffuse -in- aggregate and paratracheal Pasania formosana

24, Rays homogeneous 25

24. Rays heterogeneous 28

25. Wood semidiffuse porous 26

25. Wood diffuse porous 27

26. Parenchyma diffuse -in- aggregate, banded and paratracheal Quercus variabilis

26. Parenchyma diffuse -in- aggregate and paratracheal Quercus acutissima

27. Parenchyma apotracheal banded and Quercus phillyrae- paratracheal oides

27. Parenchyma diffuse -in- aggregate Quercus s emica rpi- and paratracheal folia

28. Tyloses absent 29 28. Tyloses present in heartwood 32 29. Growth ring indistinct 30 58

29. Growth ring distinct 31

30. Vasicentric tracheids forming large Cyclobalanopsis confluent groups pachyloma

30. Vasicentric tracheids forming Cyclobalanopsis 1- 3- seriate sheath about the vessels championii

31. Parenchyma diffuse -in- aggregate Cyclobalanopsis and paratracheal morii

31. Parenchyma apotracheal banded and C ycl obalanops is paratracheal gilva

32. Growth ring scarcely distinct 33

32. Growth ring indistinct 34

33. Parenchyma diffuse, diffuse-in- aggre- Cyclobalanopsis gate and paratracheal stenophylloides

33. Parenchyma diffuse -in- aggregate and Cyclobalanopsis paratracheal glauca

34. Parenchyma apotracheal banded and C ycl obalanops is paratracheal longinux

34. Parenchyma diffuse -in- aggregate and Cyclobalanopsis paratracheal globosa 59

DISCUSSION AND CONCLUSIONS

The foregoing description of the wood anatomy of seven gen- era and thirty five species of the Fagaceae of Taiwan indicates the complexity of the secondary xylem in this family. There is consider- able variation in the different types of xylem elements among the species studied. No single character was found to occur throughout the whole family but vasicentric tracheids are present in all genera except Fagus. All the genera can be separated easily on the basis of anatomical structures except Lithocarpus and Pasania. However, these two genera can be separated on the basis of morphology of re- productive structures. Species within each genus, especially in Pasania and Cyclobalanopsis, are in general rather difficult to separate anatomically. It is hoped that later studies of external mor- phology can be correlated with these anatomical studies to better clarify some of the taxonomic problems in this family. As Bailey

(1957) stated, "If a truly natural classification of dicotyledons is to be attained, it must be based upon harmonizing evidence from all organs and parts of the plants. It is in this synthesizing task that wood anatomy has a significant role in taxonomy."

The following descriptions attempt to characterize the wood anatomy of each genus of the Fagaceae. As indicated above, dif- ficulty was encountered with Pasania and Lithocarpus. Based on the 60 species studied, and as recognized in Taiwan, these two genera could not be separated anatomically.

Fagus (Tourn.) Linn.

Growth ring distinct. Wood diffuse porous with pores evenly distributed, polygonal; pore size transition gradual; perforations simple, reticulate and scalariform; pit cavity orbicular, oval to elliptical, aperture lenticular, tyloses present. Fibers not gelat- inous. Parenchyma apotracheal diffuse and paratracheal. Rays

homogeneous or with marginal upright cells, both compound .. and narrow. The above general description is based on the author's description of Fagus hayatae and Brown and Panshin's (1940) de- scription of Fagus grandiflora.

Castanea Tourn.

Growth ring conspicuous. Wood ring porous with pores un- evenly distributed in flame -like patches in late wood; pore size transition abrupt, pores orbicular or elliptical in early wood, poly- gonal in late wood, perforation simple or scalariform; pit cavity orbicular, aperture lenticular to elliptical; tyloses present. Vasi- centric tracheids confined to the vicinity of the larger vessels.

Fibers not gelatinous. Parenchyma diffuse -in- aggregate and para- tracheal. Rays homogeneous, uniseriate, or rarely biseriate. 61

Castanopsis Spach.

Growth ring usually distinct, rarely inconspicuous. Wood diffuse or ring porous, pores unevenly distributed in radial flame- like or stream -like clusters; pore size transition gradual or abrupt, pores orbicular in early wood, polygonal in late wood; perforation simple or scalariform; pit cavity orbicular to elliptical, aperture orbicular; tyloses present. Vascentric tracheids forming 1 -3- seriate sheath about the vessels. Fibers not gelatinous. Paren- chyma diffuse, diffuse -in- aggregate, banded and paratracheal. Rays heterogeneous, compound and narrow.

Lithocarpus B1, and Pasania Oerst.

Growth ring indistinct, scarcely distinct or distinct, Wood diffuse porous, with pores unevenly distributed in radial flame -like or stream -like clusters; pore size transition gradual, pores or- bicular to elliptical; perforation simple; pit cavity orbicular, aper- ture orbicular to lenticular; tyloses present or absent. Vasicentric tracheids forming 1- 3- seriate sheath about the vessels or confined to the vicinity of larger vessels. Fibers thin- walled, not gelatinous. Parenchyma diffuse -in- aggregate, banded and paratracheal, with

tannin or crystals. Rays heterogeneous, compound,aggregate and narrow. The above description does not include Pasania uraiana. 62

This species will be discussed in more detail later.

Quercus Linn.

Growth ring distinct or scarcely distinct. Wood semidiffuse or diffuse porous, pores scattered or unevenly distributed in radial flame -like or stream -like clusters; pore size transition gradual,

pores orbicular to elliptical; perforation simple; pit cavity orbicular

to elliptical, aperture orbicular to lenticular; tyloses present.

Vasicentric tracheids forming 1- 4- seriate sheath about the vessels. Fibers thick -walled, gelatinous. Parenchyma diffuse -in- aggregate, banded and paratracheal, with tannin or crystals. Rays homoge- neous, compound, aggregate and narrow.

Cyclobalanopsis Oerst

Growth ring indistinct, scarcely distinct or distinct. Wood diffuse porous, with pores unevenly distributed in radial flame -like or stream -like clusters; pore size transition gradual, pores orbic-

ular to elliptical; perforation simple; pit cavity orbicular, aperture lenticular or rarely orbicular; tyloses present or absent. Vasicen-

tric tracheids forming l -several- seriate sheath or large confluent groups about the vessels. Fibers thin -or thick -walled, gelatinous. Parenchyma diffuse -in- aggregate, banded and paratracheal, with

tannin or crystals. Rays heterogeneous, compound, aggregate 63 and narrow.

Key to Genera of Fagaceae in Taiwan

1. No vasicentric tracheids Fagus

1. Vasicentric tracheids present 2

2. Gelatinous fibers absent 3

2. Gelatinous fibers present 5

3. Rays homogeneous Castanea

3. Rays heterogeneous 4

4. Polygonal pores present Castanopsis

4. Polygonal pores absent Lithocarpus, Pa s ania

5. Rays homogeneous Quercus

5. Rays heterogeneous Cyclobalanopsis

According to Shimaji (1962), there are some vasicentric tracheids in Fagus, but they are scarce and restricted to occasional inconspicuous cells contiguous to vessels. However, Brown and

Panshin (1940) did not find such elements in the wood of the Ameri- can species, Fagus grandifolia, and I did not find any in Fagus hayatae. On this basis Fagus is considered to be the more primitive genus in the Fagaceae. In addition to the lack of vasicentric trache-

ids, , Fagus shows other primitive characters such as the polygonal, evenly distributed pores, the oblique perforation plates and 64 scalariform perforations.

The author has made no attempt to determine phylogenetic re- lationships among the genera of the Fagaceae. Several proposals, based on various types of evidence, suggest possible interrelation- ships of the genera in this family. Schottky (1912), on the basis of data from the leaf, the flower and cupula, concluded that the genus

Pasania is the most advanced, that the genera Castanea, Castanopsis, Quercus, Cyclobalanopsis and Pasania were derived from a common ancestor, and that divergence of the genus Fagus and the ancestor of the other genera occurred early. Schwarz (1936), on the basis of data from the flower, the cupula, the cotyledon, etc, concluded that the Quercus is the most advanced, and that all the genera evolved early and followed parallel paths in evolution. Shimaji (1962), on the basis of both external morphology and internal structure, agreed that Schottky's hypothetical relationships were reasonable but he considered that Quercus is the most advanced genus, and that Castan- opsis is more primitive than Castanea. The present study offers no new evidence regarding interrelationships among the genera of the Fagaceae, but may suggest possible solutions regarding certain species which have been variously interpreted by different authors. Usually both anatomical and morphological characters are involved in these interpretations.

Pasania uraiana is one of the most difficult species to place. 65

Originally it was described as Quercus uraiana by Hayatae (1911) who later (1916) transfered it to Lithocarpus uraiana, and it was placed in Synaedrys by Koidzumi (1916). Kanehira and Hatushima (1936)

treated it under the genus Shiia, then transfered it to the genus

Castanopsis in 1939. Masamune and Tomiya (1947) treated it as the type species of a new genus, Limlia. Recent taxonomic studies con- siderthe species in Pasania. Shimaji (1962) agreed with Kanehira and Hatushima (1939) that the species belonged in Castanopsis on the basis of wood anatomy. This does not agree with his three dimen-

sional model of phylogentic interrelationship of the groups within the

Fagaceae. According to his model, the cupula of Castanopsis is en- closed, but the cupula of Pasania uraiana is cup -like. Certainly the wood anatomy of this species is very similar to that of Castanopsis and quite different from the anatomy of Pasania or Lithocarpus. The wood resembles most closely that of Castanopsis carlesii var. ses-

silis, except that the walls of the fibers in the latter are somewhat thinner. Thus this species varies from all the species of the genus

Pasania or Lithocarpus in wood anatomical characters,. and varies from Castanopsis in external morphology. If both anatomical and morphological characters are considered, probably Masamune and

Tomiya's placement of this species in the new genus, Limlia, is the best solution. Pasania formosana and Pasania dodoniaefolia have also been 66 variously interpreted. Li (1953) treated them as one species under the name of Pasania formosana but Lin and Liu (1965) considered them to be separate and distinct species as originally de- scribed. A comparison of wood anatomy shows consistent differences between the two species. Pasania dodoniaefolia shows distinct growth rings, homogeneous rays, and absence of crystals in the parenchyma cells. Pasania formosana shows indistinct growth rings, heterogen- eous rays and crystals in parenchyma cells. Thus anatomical evi- dence supports the earlier morphological evidence used to establish these as two distinct species. Castanopsis carlesii var. sessilis was described originally by Hayata (1916) as Quercus stipitata, then transfered to Lithocarpus by Koidzumi (1925), to Shiia by Kudo and Masamune (1930), to Castan- opsis by Kanehira (1939), and finally treated as a variety of Castan- opsis carlesii by Nakai (1939), which is currently recognized by Li

(1953, 1963) and Lin and Liu (1965). Since Castanopsis carlesii has diffuse porous wood, inconspicuous growth rings, pore size trans- ition gradual, while Castanopsis carlesii var. sessilis has ring por- ous wood, conspicuous growth rings, pore size transition abrupt, these should be considered as separate species, The best solution would appear to be to. reduce 'Castanopsis carlesii var, sessilis to synony- my and recognize Castanopsis stipitata as the valid naine for this species.

Ther.e are many species in the Fagaceae that would be 67 extremely difficult to identify only on the basis of wood anatomy.

The difference between any two species is very slight. For instance, the major difference between Castanea crenata and Castanea mollis- sima is the presence or absence of scalariform perforations in the late wood; between Castanopsis formosana and Castanopsis stella- tospina, and also between Quercus variabilis and Quercus acutis- sima is only the arrangement of xylem parenchyma; between Litho - carpus amygdalifolius and Pasania nantoensis is only the presence or absence of tyloses in the heartwood; and between Pasania kawaka- mii and Pasania dodoniaefolia is only the shape of the pit aperture.

My observations differ in several minor ways from those of earlier workers on wood anatomy of the Fagaceae. Shimaji (1962) recorded the occurence of vasicentric tracheids in Fagus including Fagus hayatae and Fagus grandifolia. The tracheids were described as few and inconspicuous. Brown and Panshin (1940) did not report such elements in Fagus grandifolia and the author did not observe any in Fagus hayatae. Shimaji (1962) described the vessel distri- bution of Castanopsis hystrix as radial porous and that of Quercus acutissima as ring porous. On the basis of the samples studied by me, these woods are ring porous and semidiffuse porous, respec- tively. Other minor differences in descriptions of characteristics of various xylem elements have been found. Such variations are expected, and the smaller the sample examined, the greater will 68 be the differences reported by various workers. Carlquist (1961) has stated that there are numerous variations related to environ- ment, to location within the plant, to maturation or age of tissue, and additional variations among individuals. On the other hand, similarities in wood anatomy in different species are not indicative necessarily of close genetic relationship. This is due to the fact that parallel specialization of the tissues may have occurred inde- pendently (Bailey, 1944),

Evidence from wood anatomy of dicotyledons has proved to play an important role with reference to taxonomy and phylogeny,

The lines of specialization of the various structural features have been better established for the xylem than for any other single tissue, and have been used to clarify taxonomic affinities by many anatom- ists (Bailey, 1954; Carlquist, 1961; Metcalfe and Chalk, 1950). If the range of variation of each character of wood anatomy has been understood, and the origin of the parallel specialization of certain tissue has been found, wood anatomy will offer more significant contributions to the clarification of taxonomic affinities. 69

BIBLIOGRAPHY

1. Bailey, I. W. Contributions to plant anatomy. Waltham, Mass. , Chronica Botanica, 1954. 259 p.

2. . The development of vessels in angiosperms and its significance in morphological research. American Journal of Botany 31:421 -428. 1944.

3. . The potentialities and limitations of wood anatomy in the study of the phylogeny and classification of angiosperms. Journal of the Arnold Arboretum 38:243 -254. 1957.

4. Brown, H. P. and A. J. Panshin. Commercial timbers of the United States. New York, McGraw -Hill, 1940. 554 p.

5. Carlquist, S. Comparative plant anatomy. New York, Holt, Rinehart and Winston. 1961. 146 p.

6. Hayata, B. General index of the flora of Formosa. Taihoku, Formosa Bureau of Productive Industry, 1916. 72 p.

7. . Icone plantarum formosarum. Taihoku, Formosa Bureau of Productive Industry, 1911. 299 p.

8. Johansen, D. A. Plant microtechnique. New York, McGraw- Hill, 1940. 104 p.

9. Kanehira, R. and S. Hatushima. Formosa trees indigenous to the island. Rev. ed. Taihoku, Formosa Department of Forestry, 1936. 754 p.

10. . Additions and corrections to Formosan trees. Transactions of the Natural History Society of Formosa 29: 155 -160.. 1939.

11. Koidzumi, G. Contributions ad cognitorem florae Asiae Orientalis. XII. Botanical Magazine (Tokyo) 39:2. 1925.

12. Li, H. L. Taxonomic notes on the Fagaceae of Formosa. Bulletin of the Torrey Botanical Club 80:321. 1953. 70

13. . Woody flora of Taiwan. Norberth, Pennsylvania, Livingston Publishing Company, 1963. 974 p.

14. Lin, W. F. and T. Liu. Studies on the classification of Fagaceae in Taiwan. Taipei, 1965. 59 p. (Taiwan Forestry Research Institute. Bulletin 110).

15. Masamune, G. Three species of Formosan plants which should be included in the genus Shiia Makino. Botanical Magazine (Tokyo) 44 :405 -406. 1930. 16. Masamune, G. and Tomiya. Limlia, a new genus of Fagaceae from Formosa. Acta Botanica Taiwanica 1:1. 1947.

17. Metcalfe, C. R. and L. Chalk. Anatomy of the dicotyledons. Oxford, Clarendon, 1950. 2 vols.

18. Nakai, T. Castanopsis Pasania and their allies in the Jápan- ese Empire. Journal of Japanese Botany, 15:257 -277. 1939.

19. Schottky, E. Die Eichen des extratropischen Ostasiens und ihre Pflanzen geographische Bedeutung. Botanische Jahr - búcher 47 :617 -708. 1912.

20. Schwarz, O. Entwurf su einen natúrlichen System der Cupuliferen und der Gattung Quercus L. Notizblatt des Botanischen Gartens und Museums Zu Berlin 8:1 -22. 1936.

21. Shimaji, K. Anatomical studies on the phylogenetic inter- relationship of the genera in the Fagaceae. Bulletin of the Tokyo University Forests 57 :1 -14. 1962. APPENDIX 71

APPENDIX FIGURES

Each species is illustrated by (a) cross (b) radial and (c) tangential sections, X 60

Page

1. Fagus hayatae 73

2. Castanea crenata 73

3, Castanea mollissima 74

4. Castanopsis carlesii 74

5. Castanopsis carlesii var. sessilis 75

6. Castanopsis subacuminata 75

7. Castanopsis formosana 76

8. Castanopsis hystrix 76

9. Castanopsis kawakamii 77

10. Castanopsis kusanoi 77

11. Castanopsis stellatospina 78

12. Lithocarpus amygdalifolius 78

13. Lithocarpus castanopsisifolius 79

14. Pasania kodaihoensis 79

15. Pasania konishii - 80

16. Pasania brevicaudata 80

17. Pasania uraiana 81

18. Pasania kawakamii 81

19. Pasania dodoniaefolia 82 72

20. Pasania randaiensis 82

21. Pasania formosana 83

22. Pasania ternaticupula 83

23. Pasania nantoensis 84

24. Quercus variabilis 84

25. Quercus acutissima 85

26, Quercus phillyraeoides 85

27. Quercus semicarpifolia sub sp. glabra 86

28. Cyclobalanopsis pachyloma 86

29. Cyçlobalanopsis. morii: 87

30. Cyclobalanopsis championii 87

31. Cyclobalanopsis gilva 88

32. Cyclobalanopsis stenophylloides 88

33. Cyclobalanopsis glauca 89

34. Cyclobalanopsis longinux 89

35. Cyclobalanopsis globosa 90 73

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